©Societas Europaea Lepidopterologica; download unter http://www.biodiversitylibrary.org/ und www.zobodat.at Nota Lepi. 38(1) 2015: 89-102 DOI 10.3897/nl.38.4816 | In Memoriam: Niels Peder Kristensen (1943-2014) 13 2 1 Thomas J. Simonsen Ole Karsholt Malcolm J. Scoble , , 1 Natural History Museum, United Kingdom, London, UK 2 The Natural History Museum ofDenmark, Copenhagen, Denmark 3 Current address: Natural History Museum Aarhus, Aarhus, Denmark http://zoobank.org/F762FCAl-0D5A-4CDF-9EB3-93A 7800752FD Received 3 March 2015; accepted 18 March 2015; published: 12 May 2015 Subject Editor: Jadranka Rota. Niels Peder Kristensen, Honorary Member and former president of SEL, passed away on Satur- day December 6th 2014 in Copenhagen. While his death was not unexpected, its timing came earli- er than we had thought or hoped. His loss is felt widely and intensely. Bom on March 2nd 1943, Niels was the second child of Thorkil and Ellen Christine Kristensen (nee Nielsen). His father was an academic, politi- cian and thinker who served as Minster of Finance in two different government cabinets, and later as General Secretary of the OECD. Growing up in such an environment undoubtedly had a profound influence on Niels’ own world view, one which was powerfully international in its expression, yet retaining a strong interest and deep concern for Danish issues - local and national. Niels developed an interest in entomolo- gy and lepidopterology in particular at an ear- nd ly first time, Figure 1. Niels Peder Kristensen, March 2 1943 age, and once told TJS about the - December 6th 2014 (photo: Birgit Nielsen). when eight years old, he visited the Entomology Department at the Zoological Museum in Copen- hagen (ZMUC) ‘clutching his father’s hand’. After completing high school at Birkerod Statsskole in 1961, Niels enrolled as a biology student at the University of Copenhagen, and quickly became a regular visitor to the Entomology Department of the Museum, where he had already started as a volunteer during his last years at high school. In 1965, while still a student, he published his first paper, which was on the faunistics of Danish cicadas. From the very start of his scientific career, one of Niels’ abiding interests was the evolution (particularly evolutionary morphology) of primitive Lepidoptera. Indeed, the work for his Mag. Scient, degree was on the comparative morphology of the primitive glossatan family, Eriocraniidae. During this study, Niels spent the academic year 1966-67 at the University of Bristol, working with the eminent and extremely knowledgeable Brit- ©Societas Europaea Lepidopterologica; download unter http://www.biodiversitylibrary.org/ und www.zobodat.at 90 SiMONSEN et al: In Memoriam: Niels Peder Kristensen (1943-2014) ish entomologist Howard E. Hinton, who was at that time pioneering the use of scanning electron microscopy in entomology. It was while working with Hinton that Niels came to appreciate the value of SEM in comparative morphology. Through it, he made the surprising discovery (published in 1970) that the most primitive Lepidoptera have flat, solid wing scales (i.e. lacking an internal lu- men), a condition contrasting strongly with the hollow wing scales generally found in Lepidoptera. After obtaining his Mag. Scient, degree in 1968, Niels was offered a tenure-track position at the ZMUC as Amanuensis (Assistant Professor). He was promoted to Associate Professor in 1972 and appointed as Full Professor of Entomology in 1995. In 1970, Niels visited one of Europe’s foremost comparative invertebrate morphologists, Jean Chaudonneret, at the Université de Dijon where he enhanced considerably his skills in insect histology and semi-thin sectioning. Working with Hinton, Chaudonneret and Karl G. Wingstrand, the professor of comparative morphology at the University of Copenhagen, unquestionably had a profound influence on Niels’ development as a scientist. He often referred to the effect these three mentors had on his career. Niels was also deeply interested in the analytical methods used in evolutionary research. Together with fellow entomologist Nils Möller Andersen and the palaeontologist Niels Bonde, he was a pioneer in Den- mark, and more widely in Scandinavia, of Hennig’s phylogenetic systematics, and his cladistic analyses of the higher-level relationships of butterflies in 1976 remained the standard work on the subject until the study by de Jong et al. (1996) 20 years later. From the very start of his career, Niels was deeply interested in the morphology and phylogeny of the higher insects. In 1975 he published (Z. zool. Syst. Evolut. -forsch. 13, pp. 1^14) one of his most influential papers: “The phylogeny of hexapod ‘orders’. A critical review of recent accounts”. Thirty years later, Grimaldi and Engel (2005, p. 144) referred to this work as “perhaps the single most important paper in systematic entomology”. This publication formed the basis of his five updated reviews of the subject. The last of these was published in Eur. J. Ent. in 1999, while perhaps the most notable of them is the 1991 text-book chapter “Phylogeny of extant hexapods” in “The Insects of Australia”, which should be mandatory reading for all students of systematic entomology. Over the years Niels authored or co-authored a number of papers on higher Hexapod relationships especially on the lower Hexapod orders, Trichoptera, the enigmatic New Zealand mecopteran family Nanno- choristidae, Neuroptera, and of course Mantophasmatodea—the first new insect order to be described for 90 years, the description of which he co-authored in 2004. It was, however, the Lepidoptera that remained Niels’ main interest, and the majority of his publi- cations are on that order. They range in scope from nomenclatural and faunistic notes to higher-lev- el phylogenetics and to the exceptionally detailed, comparative morphological studies of primitive Lepidoptera. More than anything else, these exquisite studies became his professional hallmark. His early enthusiasm for scanning electron microscopy and histology were combined with transmission electron microscopy and became methodological cornerstones in his work throughout his working life. Much of his productivity, particularly in the first half of his career, led to highly detailed studies of lit- tle-understood structures and organ systems of primitive Lepidoptera, including overall head and neck anatomy, mouthpart morphology, anatomy of the alimentary canal, structure of the trachaea system, comparative morphology and anatomy of male and female genitalia, and wing scales and vestiture. Niels’ work on primitive Lepidoptera morphology and anatomy was always embedded in the context of higher Lepidoptera evolution, and his ultimate goal was to establish the early evolutionary patterns within the order, thereby creating a sound basis for further studies higher up the lepidopteran tree. In 1978 and 1979 he also described two new families of primitive Lepidoptera, the basal hepialoid family ©Societas Europaea Lepidopterologica; download unter http://www.biodiversitylibrary.org/ und www.zobodat.at Nota Lepi. 38(1): 89-102 91 Figure 2. Niels studying his beloved homoneuran Lepidoptera (photo: TJS). Neotheoridae and the non-ditrysian family Heterobathmiidae (the latter in collaboration with the late Ebbe S. Nielsen). His work on primitive Lepidoptera phylogeny and comparative morphology culmi- nated in his Dr Scient, dissertation “Studies on the morphology and systematics of primitive Lepidop- tera” published in Steenstrupia in 1984. Until the modifications introduced by the very recent advent of phylogenomic studies and especially the surprising discovery of a new primitive moth family from Australia, this remained the standard work on the evolution of the homoneurous Lepidoptera. In the early 1990s Niels was appointed the editor-in-chief of the two Lepidoptera volumes of the Handbook of Zoology. This immense undertaking was to dominate his professional life for the following decade. The two volumes, which were published in 1998 and 2003, defined the latter part of his career as much as his work on higher Hexapod phylogeny and comparative Lepidoptera mor- phology had shaped his early and mid career, although he continued his work on these topics until illness forced him to stop just weeks before his death. Niels had anticipated writing or co-authoring a substantive part of the first volume. He did not, however, expect to have made a similar input to the second volume, which was on morphology and physiology. Having to do so resulted in a much greater effort on his part than he had intended: moreover, it required him to write about subjects on which he did not consider himself an expert. The result, nevertheless, stands as a landmark publi- cation and a tribute to Niels’ capacity and breadth of knowledge. The Handbook would have been more than enough of a mega-project for most of us, so it is remarkable that Niels also spent much time and effort during his last years editing a book on the insects of Greenland instead of completing some of his own research projects. While he certainly believed in the value of the Greenland work, his resolve was propelled by that innate sense of responsibility and conscientiousness that were so evident in his personal makeup. ©Societas Europaea Lepidopterologica; download unter http://www.biodiversitylibrary.org/ und www.zobodat.at 92 SiMONSEN et al: In Memoriam: Niels Peder Kristensen (1943-2014) After the mammoth task of completing the Handbook, Niels returned to his work on primitive